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UNIVERSIDAD DE COSTA RICA SISTEMA DE ESTUDIOS DE POSGRADO ESTUDIO MORFOLÓGICO, FILOGENÉTICO Y FENOLÓGICO DE Smilax L. (SMILAC

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UNIVERSIDAD DE COSTA RICA SISTEMA DE ESTUDIOS DE POSGRADO ESTUDIO MORFOLÓGICO, FILOGENÉTICO Y FENOLÓGICO DE Smilax L. (SMILAC UNIVERSIDAD DE COSTA RICA SISTEMA DE ESTUDIOS DE POSGRADO ESTUDIO MORFOLÓGICO, FILOGENÉTICO Y FENOLÓGICO DE Smilax L. (SMILACACEAE) EN COSTA RICA, CON IMPLICACIONES SISTEMÁTICA~ Tesis sometida a la consideración de la Comisión del Programa de Estudios de Posgrado en la Escuela de Biología para optar al grado de Magíster Scientiae LILIAN FERRUFINO ACOST A Ciudad Universitaria "Rodrigo Facio", Costa Rica 2003 DEDICATORIA A MIS PADRES CECILO FERRUFINO Y TERESA ACOSTA A MI NOVIO GREGORIO DAUPHIN POR SU APOYO Y COMPRENSIÓN AGRADECIMIENTOS Deseo agradecer a las instituciones y personas que con su apoyo hicieron posible esta investigación A Dios quien siempre escucha mis oraciones y plegarias cada momento Al Servicio Alemán de Intercambio Académico (DAAD), por otorgarme la beca a través del programa regional para cubrir mis estudios de posgrado. A mi comité de tesis: al M. Se. Jorge Gómez-Laurito, al Dr. Osear Rocha Núñez y al Dr. Carlos O. Morales por su apoyo y asesoramiento. Al Jardín Botánico de Missouri que a través de las becas Elizabeth Bascom me apoyaron en la revisión de literatura y examinación de especímenes tipos. Al personal científico del Jardín Botánico de Missouri, en especial a la M. en C. Alina Freire­ Fiero y al Dr. Gerrit Davidse por sus comentarios en el manuscrito del tratamiento taxonómico. Al Centro Agronómico Tropical de Investigación y Enseñanza (CATIE), al Instituto Tecnológico de Costa Rica sede en San Carlos, a la Finca Coope San Juan y al Jardín Agroecológico Bougainvillea por su apoyo económico y logístico. A mis amigos por su apoyo moral y logístico y en especial a Gregorio Dauphin. Al personal del laboratorio de genética de la Escuela de Biología, en especial a Gabriel Aguilar por su asesoramiento en la parte práctica de sistemática molecular. di HOJA DE APROBACIÓN Esta Tesis fue aceptada por la Comisión del Programa de Estudios de Posgrado en Biología de la Universidad de Costa Rica, como requisito parcial para optar al grado Magíster Scientiae M. Se. Elmer García Díaz Representante de la Decana del Sep M. Se. Jorge Gómez-Laurito Director de Tesis Dr. Osear J. Rocha Núñez Asesor Dr. Carlos Morales Sánchez Asesor Dra. Virginia Solís Alvarado Directora del Programa de Posgrado en Biología Lilian F errufino Acosta Sustentante PREFACIO Esta Tesis fue escrita de acuerdo con el Reglamento del Sistema de Estudios de Posgrado en Biología de la Universidad de Costa Rica. Está dividida en tres capítulos y cada uno sigue las reglas de presentación de manuscritos de la Revista de Biología Tropical. INDICE GENERAL DEDICATORIA .............................................................................................................................................. ii AGRADECIMIENTOS .................................................................................................................................. iii HOJA DE APROBACIÓN ............................................................................................................................. h· PREFACIO ...................................................................................................................................................... v INDICE GENERAL ....................................................................................................................................... vi LISTA DE FIGURAS .................................................................................................................................... vii LISTA DE CUADROS···································································································································· X LISTA DE ANEXO......................................................................................................................................... xi RESUMEN ..................................................................................................................................................... xii ARTÍCULO l. ESTUDIO MORFOLÓGICO DE Smilax L. (SMILACACEAE) EN COSTA RICA, CON IMPLICACIONES SISTEMÁTICAS.................................................................................•........................... 1 ARTICULO II. FILOGENIA DE Smila.x L. (SMILACACEAE) EN MESOAMÉRICA............................ 70 ARTICULO 111. ESTUDIO FENOLÓGICO DE Smilax L. (SMILACACEAE) EN COSTA RICA. ......... 86 \I LISTA DE FIGURAS ARTÍCULO 1 FIGURA# CONTENIDO PÁGINA 1 Localización de las tres áreas de estudio en Costa Rica. 33 2 Granos de polen de Smilax. A, B, C, D, E. Vista general; F. 40 Detalle de la exina con gránulos y espínulas. A S. domingensis; B. S. mol/is; C. S. spinosa; D y F. S. subpubescens; E. S. vanilliodora. 3 Smilax domingensis Willd.: A Flor femenina, B. Flor 47 masculina, C. Estambre, D. Frutos, E. Semillas, F. Rama fértil con agallas y flores, G. Tallo y rizoma. 4 Distribución geográfica de Smilax domingensis ( •) y S. 51 mollis (~) en Costa Rica. 5 Smilax mollis Humb. & Bonpl.: A Flor femenina, B. Flor 55 masculina, C. Estambre, D. Frutos, E. Semillas, F. Rama joven, G. Rama fértil con agallas y botones, H. Tallo y rizoma. 6 Smilax panamensis Morong: A Flor femenina, B. Flor 59 masculina, C. Estambre, D. Semillas, E. Rama fértil con fiutos, F. Tallo y rizoma. 7 Distribución geográfica de Smilax panamensis ( •) y S. 61 spinosa (~) en Costa Rica. 8 Smilax spinosa Mill.: A Frutos, B. Semillas, C. Flor 65 femenina, D. Flor masculina, E. Estambre, F. Rama fértil con flores, G. Tallo, H. Tallo y rizoma. 9 Smilax spissa Killip & C. Morton: A Flor femenina, B. Flor 68 masculina, C. Estambre, D. Semillas, E. Rama fértil con frutos, F. Tallo y rizoma 10 Distribución geográfica de Smilax ~pissa ( •) y S. 69 subpubescens (~)en Costa Rica. FIGURA# CONTENIDO PÁGINA 11 Smilax subpubescens A. DC.: A. Flor femenina, B. Flor 71 masculina, C. Estambre, D. Frutos, E. Semillas, F. Rama fértil con flores, G. Tricomas de la base del peciolo, H. Tallo y rizoma. 12 Smilax vanilliodora Apt A. Flor femenina, B. Flor 78 masculina, C. Estambre, D. Semillas, E. Rama fértil con frutos, F. Tallo y rizoma. 13 Distribución geográfica de Smilax vanilliodora ( •) en Costa 80 Rica. ARTÍCULO 11 FIGURA# CONTENIDO PÁGINA Figura 1. Arbol de consenso estricto de Smilax (Sección 99 Micranthae) en Mesoamérica, basado en caracteres morfo lógicos. 2 Figura 2. Árbol de consenso de Jacknife para las especies de 100 Smilax (sección Micranthae) basado en caracteres morfológicos. 3 Figura 3. Árbol de consenso representando el número de 100 caracteres para cada clado. El número del carácter corresponde al nombre del carácter que se encuentra en el Cuadro l. 0 = homoplasias, • = sinapomorfias. ARTÍCULO 111 FIGURA# CONTENIDO PÁGINA l Localización de las tres áreas de estudio en Costa Rica. 114 2 Promedios de caída de hojas de cinco especies de Smilax en 116 Costa Rica. A. Reserva Biológica Alberto Manuel Brenes (RBAMB), B. Finca Coope San Juan, y C. Jardín Agroecológico Bougainvillea (JAB). \'111 FIGURA# CONTENIDO PÁGINA 119 3 Promedios de brotadura de hojas de cinco especies de Smilax en Costa Rica. A Reserva Biológica Alberto Manuel Brenes (RBAMB), B. Finca Coope San Juan, y C. Jardín Agroecológico Bougainvillea (JAB). 4 Promedios de floración de Smilax en Costa Rica. A Reserva 122 Biológica Alberto. Manuel Brenes (RBAMB) y B. Jardín Agroecológico Bougainvillea (JAB). 5 Promedios de fructificación de Smilax en tres sitios en Costa 124 Rica. A Reserva Biológica Alberto Manuel Brenes (RBAMB), B. Jardín Agroecológico Bougainvillea (JAB) y C. Finca Coope San Juan. IX LISTA DE CUADROS ARTÍCULO 11 CUADRO# CONTENIDO PÁGINA 1 Caracteres y estados de caracteres morfológicos usados en 97 el análisis filogenético de Smilax (Sección Micranthae). ARTÍCULO 111 CUADROS# CONTENIDO PÁGINA 1 Análisis de varianza (SAS Inc., 1999-2000) para los 117 eventos fenológicos producidos en la Reserva Biológica Alberto Manuel Brenes (RBAMB) en San Ramón, la Finca Coope San Juan, y el Jardín Agroecológico Bougainvillea. 2 Prueba Sheffe para cinco especies de Smilax en Costa 117 Rica. Entre paréntesis el promedio. 3 Correlación de Pearson de las especies de Smilax 120 estudiadas en tres sitios. LISTA DE ANEXO ARTÍCULO U ANEXO CONTENIDO PÁGINA 1 Especímenes y fuente del material vegetal usado en la 106 extracción y secuenciación de ADN de la región intergénica tmL-trnF. ESTUDIO MORFOLÓGICO, FILOGENÉTICO Y FENOLÓGICO DE Smilax L. (Smilacaceae) EN COSTA RICA, CON IMPLICACIONES SISTEMÁTICAS F errufino A costa, Lilian Florencia Estudio morfológico, filogenético y fenológico de Smilax L. (Smilacaceae) en Costa Rica, con implicaciones sistemáticas Tesis Magister Scientiae en Biología. San José, C. R.: L. F. Ferrufino A, 2003 l 13h.: 21 iL- 200 refs. RESUMEN INTRODUCCIÓN Taxonomía Smilax consta aproximadamente de 300 especies en zonas templadas y tropicales en ambos hemisferios (Judd et al. 2002). Las plantas de Smílax son bejucos subleñosos, rara vez herbáceos, dioicos, que se originan de un rizoma delgado o grueso. Usualmente trepan mediante pares de zarcillos que se originan en los pecíolos y a menudo poseen aguijones en tallos y hojas. Poseen hojas alternas con 3 a 9 nervios paralelos o triplinervias, conectados por nervaduras reticuladas. Tienen flores pequeñas, dispuestas en umbelas axilares, a veces racemosas, con 6 tépalos libres, iguales, 6 estambres libres, ovario súpero, 1 ó 2 óvulos en cada lóculo; estaminodios presentes en las flores femeninas. Los frutos son bayas globosas, negras, púrpuras, rojas o anaranjadas (Cronquist 1981, Huft 1994, Gómez-Laurito 1998). Se considera que en Mesoamérica existen
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